Новости Сист. Низш. Раст. 51 — Novosti Sist. Nizsh. Rast. 51
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Новости сист. низш. раст. — Novosti Sist. Nizsh. Rast. 51: 242–250. 2017 Gemmae in Tetralophozia setiformis (Anastrophyllaceae, Marchantiophyta) and their second record in Eurasia M. V. Dulin Institute of Biology, Komi Science Centre UB RAS, Kommunisticheskaja Str., 28, Syktyvkar, Komi Republic, 167982, Russia; [email protected] Abstract. Tetralophozia setiformis is a widespread species occurring usually with- out organs of sexual and asexual reproduction. Gemmae of Tetralophozia setiformis were observed for the second time in Russia and Eurasia in the Northern Urals, Komi Republic. They form compact masses over upper leaves. The compact masses consist largely (70 %) of immature gemmae. Description of gemmae and gemmiparous shoots from the Northern Urals and their comparison with those from the other known locali- ties, namely British Columbia (Canada) and the Murmansk Region (European Russia) were carried out. The gemmiparous plants of T. setiformis from the Northern Urals have approximately the same width as plants without gemmae but they are shorter. The leaves of gemmiparous plants from the Northern Urals are similar to leaves of gemmiparous plants from British Columbia. The leaf shape in upper part of the gem- miparous shoots varies from the typical to ± modifi ed from gemmae production. These leaf shape transitions include reduction of leaf size and lobe number from 4 to 2–3, suppression of development and disappearance of characteristic teeth at the base of sinus. Gemmae size (17 × 22 μm) of plants from the Northern Urals is within vari- ability recorded for plants from the Murmansk Region and British Columbia. Keywords: gemmae, liverworts, Tetralophozia setiformis, British Columbia, Mur- mansk Region, Northern Urals, Komi Republic, Russia. Выводковые почки у Tetralophozia setiformis (Anastrophyllaceae, Marchantiophyta) и их вторая находка в Евразии М. В. Дулин Институт биологии Коми НЦ УрО РАН, ул. Коммунистическая, д. 28, г. Сыктывкар, 167982, Россия; [email protected] Резюме. Tetralophozia setifomis — широко распространенный вид, обычно встречающийся в вегетативном состоянии без органов полового и бесполого размножения. Описана вторая находка выводковых почек T. setifomis в России и Евразии и первая на Северном Урале, в Республике Коми. Приведено описа- ние найденных на Северном Урале выводковых почек, побегов, образующих выводковые почки, и их сравнение с данными из двух других известных место- нахождений в Британской Колумбии (Канада) и Мурманской области (Россия). Почкующиеся побеги приблизительно такие же по ширине, но более короткие, с выводковыми почками на верхних листьях в плотных скоплениях, которые об- разованы преимущественно (на 70 %) недозрелыми выводковыми почками. Ли- стья побегов с выводковыми почками с Северного Урала сходны с таковыми из Британской Колумбии. Описаны переходы формы листьев с верхушек почкую- 242 Dulin. Gemmae in Tetralophozia setiformis щихся побегов от типичных до формирующих выводковые почки, у которых на- блюдалось наряду с уменьшением размеров уменьшение числа лопастей от 4 до 2–3 также уменьшение и исчезновение характерных зубцов в основании выре- зок. Выводковые почки растений с Северного Урала по размерам (17 × 22 мкм) соответствуют ранее выявленному диапазону изменчивости этого признака, но максимальная длина у них меньше. Ключевые слова: выводковые почки, печеночники, Tetralophozia setifomis, Британская Колумбия, Мурманская область, Северный Урал, Республика Коми, Россия. The genus Tetralophozia (R. M. Schust.) Schljakov of the fami- ly Anastrophyllaceae L. Söderstr., De Roo et Hedd. includes four spe- cies known from Arctic to tropical latitudes in Asia and Africa (Schus- ter, 2002; Söderström et al., 2010; Söderström et al., 2016). The type of the genus, an arctic-alpine imperfectly circumpolar species T. setiformis (Ehrh.) Schljakov [≡ Chandonanthus setiformis (Ehrh.) Lindb.] is most widespread in the Northern Hemisphere (Konstantinova, 2000; Damsholt, 2002; Schumacker, Váňa, 2005; Konstantinova et al., 2009). Of the four species currently recognized in the genus Tetralophozia (Söderström et al., 2016), gemmae are known only in this species (Schuster, 2002). The plants of T. setiformis are fragile when dry and usually dissemi- nate via fragments. They produce sporophytes and gemmae very rarely. The gemmae were fi rst recognized by Godfrey and Godfrey (1978) from W. B. Schofi eld’s collections obtained from British Columbia. In recent European liverwort identifi cation books (Schljakov, 1980; Damsholt, 2002) there are no description of gemmae. Konstantinova (2002) in her treatment of the genus Tetralophozia for Russia briefl y described and il- lustrated gemmae of T. setiformis for the fi rst time for Eurasia from the Murmansk Region, Northwest European Russia, however, this reference tends to be overlooked. Then, Damsholt (2013) mentioned that gemmae in T. setiformis are known only from western Canada. In 2016 gemmae of T. setiformis were observed in a collection from the Northern Urals (Komi Republic) that represent the second record of the gemmae in T. setiformis in Eurasia. The purpose of the present paper is to summarize known data on the gemmae of T. setiformis and to provide comparisons from three known distant localities, illustrations and analysis of conditions of their formation. Materials and Methods Habitats and reproduction of Tetralophozia setiformis in the Northern Urals. In the Northern Urals, T. setiformis grows on and be- tween stones at stony fi elds, as well as in small holes and cracks with fi ne 243 Новости сист. низш. раст. — Novosti Sist. Nizsh. Rast. 51: 242–250. 2017 earth in rocky outcrops (remnants and cliffs). It rarely occurs on stones in dwarf-shrub and lichen tundra and on stones along mountain streams. It occurs also on stones in upland light forests, fi r-spruce forests and bushes of Betula nana L. The species occurs in large compact patches or tufts, usually with greenish-, yellowish- or golden-brown to dark brown pig- mentation or as single shoots among other bryophytes such as Tritomaria quinquedentata (Huds.) H. Buch, Sphenolobus minutus (Schreb.) Berggr. Many populations of T. setiformis were observed earlier in the Komi Re- public without organs of sexual or asexual reproduction and reproduce apparently by parts of the shoots and leaf fragments. Plants of T. setifor- mis are very fragile during dry periods and disseminated by wind, ani- mals, etc. Location of gemmiparous plants of Tetralophozia setiformis in the Northern Urals. Russia: Komi Republic, Northern Urals, Vuktylsky Dis- trict, «Yugyd Va» National Park, the Shchugor River basin, near Halmer- sale Mountain, 3 km SWW of the merger of sources of Halmerya River, 63°48ʹ56.8ʺN; 59°10ʹ48.5ʺE, ca. 715 m alt., lake cirque, steep (45–60°) slope of the cirque, NE exposure, rock outcrops, on ledge in yellowish- brown patches with whitish patches of degraded shoots, 01.07.2016 M. V. Dulin 3ha=1237mvd, SYKO. Gemmiparous shoots were studied using microscopes LOMO MBS 10 and Micmed-2. Shoots were measured after moistening. Prepared leaves and gemmae were selected in separate specimen fi xed by glycerin after microscopy. All observed gemmae (80) and 22 leaves were measured and kept after investigation as separate slides. Maximum values and mean values were calculated on the basis of the obtained measurements. Results Only two gemmiparous shoots of T. setiformis were observed in the specimen cited above. It should be noted that the specimen looks atypical. It contains the plants growing parallel to each other and assembled in compact patches with shoots spread in different directions and interwoven. The latter shoots are shorter and curved. Besides, the main part of plants (up to 50 %) in the specimen has features of degradation expressed in partial or complete yellowing of the plants. It is diffi cult to understand the reasons of such degradation. But, possibly, one of the reasons is being adjacent to the lichen Cladonia coccinifera (L.) Willd. that may affect the plants of T. setiformis by releasing lichen acids characterized by a high antibiotic activity. Such unfavorable condition resulted in degradation of T. setiformis may have initiated gemmae formation in the studied plants. 244 Dulin. Gemmae in Tetralophozia setiformis Description of gemmiparous plants and gemmae of Tetralophozia setiformis from the Northern Urals (Plate I). The gemmiparous plants of T. setiformis are golden-green to brown. They are smaller in size than non-gemmiparous plants from the same specimen (Table). At the old part of shoots they are 590–760 μm wide, in upper younger narrowed part of the shoots — 320 μm wide and 0.8–1.0 cm long. The leaves of gemmiparous shoots are divided for 0.7–0.9 their length into 3–4 lobes, × 1.0–1.5 as wide as long, 430–700 μm wide × 420–630 μm long; middle leaf lobes are × 1.2–1.9 as long as wide, 310–350 μm long × 160–270 μm wide. At the younger narrowed part of shoots, leaves × 1.0–1.1 as wide as long, 270–320 μm wide × 270–290 long; middle leaf lobes × 1.7–1.8 as long as wide, 170–180 μm long × 100–110 μm wide. Table Variability of morphological features of Tetralophozia setiformis in collections with gemmae Regions Features Murmansk Northern Urals British Columbia Region Nongemmiparous 0.6–1.5 to 6.0 1.6–4.0 shoot length, cm Nongemmiparous (430)540–590(760) (500)700–1100 (680)760–975 shoot width, mm Nongemmiparous (380)430–790(860) to 1000 (770)825–1075 leaf width, μm Nongemmiparous (320)380–540 400–600 (425)450–600 leaf length, μm Nongemmiparous 1.2–1.6 1.7 1.6–2.0 leaf, width/length ratio Gemmiparous shoot 0.8–1.0 «small», no 1.5–2.6 length, cm length mentioned Gemmiparous shoot (320) 590–760 «small», no 400–600(760) width, mm width mentioned Gemmiparous leaf (270–320)430–700 — 350–550 width, μm Gemmiparous leaf (270–290)420–630 — 275–420 length, μm Gemmiparous leaf, 1.0–1.5 — 1.1–1.5 width/length ratio Gemmae size, μm mature: (10)13–18(23) × mature 17–20 × 10–26 × 16–44 (13)18–23(31); 20–30(45) immature: (13)16–18 × (16)18–26(33); juvenile: (13)16–18(21) × (16)18–26(31) 245 Новости сист.